243808 The South Africa Ophiocomidae) of

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Western Indian Ocean J Mar. Sci. Vol. 10, No. 2, pp. 137-154, 2012
© 2012 WIOMSA
243808
The Ophiocoma species (Ophiurida: Ophiocomidae) of
South Africa
Jennifer M. Olbers 1 and Yves Samyn 2
1
Zoology Department, University of Cape Town, Private Bag X3, Rondebosch, 7701, South
Africa; 2Belgian Focal Point to the Global Taxonomy Initiative, Royal Belgian Institute of
Natural Sciences, Belgium.
Keywords: Ophiocoma, neotype, taxonomy, Ophiuroidea, Indo-West Pacific,
Western Indian Ocean, KwaZulu-Natal.
Abstract-This study raises the number of Ophiocoma species recorded in South
Africa from four to eight. All species are briefly discussed in terms of taxonomy,
geographic distribution and ecology. In addition, the juvenile of 0. brevipes, found
on the underside of adult Ophiocoma brevipes specimens, is described in detail. A
neotype is designated for 0. scolopendrina.
INTRODUCTION
The circumtropical family Ophiocomidae
holds some of the more dominant and
conspicuous ophiuroid species present on
coral and rocky reefs. The family is rich, with
eight genera, two of which were relatively
recently reviewed by Devaney (1968; 1970;
1978). One of these, Ophiocoma Agassiz,
1836, is well represented in the tropical to
subtropical waters of KwaZulu-Natal in
South Africa and its constituent species are
documented here.
Ophiocoma species are difficult to identify
since some of the distinctive taxonomic
characters, such as the shape of the oral and
dental plates and their associated papillae (see
Devaney 1970), are not easily assessed or can
change during growth, as is also the case for
the number of arm spines, the disc armament
and the size of the dorsal arm plates (Sumida
et al. 1998; StOhr 2005; StOhr et al. 2008).
In addition, some species can change their
colouration from day to night (Hendler 1984).
Corresponding author: JMO
E-mail: [email protected]
The
Indo-Pacific
distribution
of
Ophiocoma has been dealt with by several
authors (e.g. Clark & Rowe 1971; Cherbonnier
& Guille 1978; Rowe & Gates 1995). Clark
and Rowe (1971) listed 11 species from the
Indo-West Pacific (including the Red Sea
and the Persian Gulf). Since then, a few new
species have been added (Rowe & Pawson
1977; Bussarawit & Rowe 1985; Soliman
1991; Benavides-Serrato & O'Hara 2008),
bringing the total number of valid species in
the Indo-Pacific to 13 1• The ophiuroid fauna
of southern Africa have been studied by many
specialists (Bell 1905; 1909; Clark 1923;
Mortensen 1925; 1933; Balinsky 1957; 1969;
Clark 1974; 1977; 1980; Clark & Courtman1
Ophiocoma anaglyptica Ely; 0. aegyptiaca Soliman,
1991, 0. brevipes Peters, 0. cynthiae BennavidesSerrato & O'Hara, 0. dentata Muller & Troschel,
0. endeani Rowe & Pawson, 0. erinaceus Muller
& Troschel, 0. occidentalis Clark, 0. pica Milller &
Troschel, 0. pus ilia (Brock), 0. schoenleinii Milller &
Troschel, 0. scolopendrina (Lamarck), 0. valenciae
Muller & Troschel.
JENNIFER M. OLBERS and YVES SAMYN
138
Stock 1976) but, since
Clark
and
CourtrnanStock's (1976) work, no
Ophiocoma species have
been added to the fauna of
southemAfrica. Samynand
Thandar (2003), however,
mention that de novo
sampling along the coast of
KwaZulu-Natal revealed
new
ophiuroid
many
records.
Unfortunately,
the latter authors did not
include their species list.
Until now, only four
species of Ophiocoma
have been recorded in
South Africa (Clark &
Courtman-Stock
1976):
0. erinaceus Muller &
Troschel 1842, 0. pica
Muller & Troschell842, 0.
scolopendrina (Lamarck
1816) and 0. valenciae
Muller & Troschel 1842.
Recent sampling along the
north-east coast of South
Africa (see also Samyn &
Thandar 2003), yielded
four additional Ophiocoma
species. Moreover, we
discovered an undocumented association between individuals
which we believe to be adult and juvenile 0. brevipes.
Abbreviations:
A.L.
Ann length
Disc diameter
KwaZulu-Natal
KZN
MHNG Museum d 'Historie Naturelle, Geneve, Switzerland
MNHN Museum national d'Historie Naturelle, Paris, France
RMCA Royal Museum for Central Africa, Tervuren, Belgium
Natural History Museum, London, United Kingdom
NHM
iZiko South African Museum, Cape Town, South Africa
SAM
D.D.
ZMB
Museum fiir Naturlrunde an der Universitat Humboldt m Berlin, Berlin, Germany
MATERIALS and METHODS
Specimens were collected by hand in the intertidal zone and by
SCUBA diving up to 32 m depth, during six expeditions (August
1999, July 2000, February 2001, July 2003, January 2010 and
October 2010) at several localities along the coast ofKZN, South
Africa (Fig. 1).
Specimens were anaesthetized by placing them in freshwater
and gently manipulating them so that their arms were extended
as far as possible until they ceased moving. Thereafter, they were
put in 100 %buffered alcohol for one day and transferred to 70 %
buffered alcohol for transport to the laboratory where they were
dried for permanent storage. The specimens were photographed
after preservation.
Specimens are deposited in the collections of the RMCA(Belgium)
and in the SAM (South Africa). These collections were examined
together with other vouchers in the RMCA (e.g. from Kenya, the
Seychelles and Inhaca; see also Clark 1980; 1984) as well as some
ophiocomids from the Indo-Pacific Ocean in the MNHN (France).
"'I"
"1".
a:~
•I" ~ "1"
"'f'
"'I"
,
"1"
...
"1'
-~.
,.,.
"f'
......
KOSI Bay
Bhan ga Nek
...,..
.,.,,.
,..,..
.,....
.
.,
Umhlalt
Durban
/"'
.,,..
AltWal Shoal
ar1<R)'Tl.,
7no
~'-.. •.!.,..,_,-.;
r..
~$
ortSt Johns
Umgazana
olora
..:.. ,...
,.,.
,;,
90
180
...:.. """' , ...
Figure 1. Sites where Ophiocoma species have been recorded in South Africa.
270
li'.O
,..,.,
360 Kllomoters
....
,.:..,
,..,.
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
139
Table 1. Records of the present South African collection of Ophiocoma species, showing
location, depth, museum collection numbers and the number of specimens.
Valid species name
Location
Ophiocoma brevipes
Aliwal Shoal
Sodwana Bay
20
14
Sodwana Bay (2 mile reef)
15
15
11
Sodwana Bay (Mbibi)
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Sodwana Bay (9 mile reef)
Sodwana Bay (9 mile reef)
Sodwana Bay (2 mile reef)
Sodwana Bay (Mabibi)
Aliwal Shoal
Sodwana Bay (Diep Gat)
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Aliwal Shoal
Sodwana Bay
Bhanga Nek
Aliwal Shoal
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Depth (m)
11
6
15
16
13
10
11
15
14
13
Collection number (no. of specimens)
RMCAMT2199(1)
RMCAMT 2207 (1)
RMCAMT2175 (1)
RMCAMT2215(1)
RMCAMT2191 (1)
RMCA MT 2157 (1)
RMCA MT2192 (1)
RMCA MT 2225 (1)
RMCAMT2222 (1)
RMCAMT2194 (1)
RMCAMT2195 (1)
RMCAMT2198(2)
RMCAMT2179(1)
RMCAMT2212(1)
RMCAMT2193 (1)
RMCA MT2148 (3)
RMCA MT 2208 (2)
RMCA MT 2203B1S (2)
RMCAMT2!99 (I)
SAMA28111 (1)
RMCA MT2341 (2, Juvenile)
SAM A28112 (2, Juvenile)
Sodwana Bay (2 mile reef)
16
20
14
15
14
Ophiocoma cf dental a
Sodwana Bay
20
RMCAMT2380(1)
Ophiocoma doederleini
Bhanga Nek (Saxon Reef)
Sodwana Bay (2 mile reef)
Bhanga Nek
20
15
16
RMCA MT 2249 (1)
RMCAMT2250(1)
RMCA MT 2203B1S (2)
Ophiocoma erinaceus
Sodwana Bay (5 mile reef)
Bhanga Nek (Saxon Reef)
Sodwana Bay (2 mile reef)
Bhanga Nek
Sodwana Bay (1/4 mile reef)
Sodwana Bay (7 mile reef)
18
20
15
RMCAMT2136(3)
RMCAMT2142 (2)
RMCAMT2253 (1)
RMCA MT 2155 (!,juvenile)
RMCAMT 2220 (!,juvenile)
RMCAMT2349(1)
Ophiocoma pica
Qolora, Eastern Cape
Ophiocoma pus ilia
Aliwal Shoal
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Bhanga Nek
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Aliwal Shoal
Sodwana Bay (Deep sponge)
Aliwal Shoal
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Sodwana Bay (2 mile reef)
Ophiocoma scolopendrina
Umgazana, Eastern Cape
Ophiocoma valenciae
ParkRynie
Park Rynie
Sodwana Bay (5 mile reef)
Sodwana Bay (5 mile reef)
Sodwana Bay (5 mile reef)
Aliwal Shoal
Aliwal Shoal
Aliwal Shoal
Aliwal Shoal
Aliwal Shoal
Durban
Durban
Port St Johns
Umhlali
20
10
17
Intertidal
20
13
l3
15
20
15
14
16
32
13
10
10
14
SAM A 23248 (2)
RMCA MT2345 (1)
RMCA MT2219 (I)
RMCAMT2200 (I)
RMCAMT2346 (I)
RMCAMT2337 (I)
RMCAMT2217 (1)
RMCA MT 2214 (I)
RMCA MT 2218 (I)
RMCAMT2380(1)
RMCA MT 2153 (1)
RMCAMT2221 (1)
RMCAMT2149(1)
RMCAMT2381 (I)
SAM A 23093 (2)
Intertidal
Intertidal
18
16
18
16
13
13
13
13
Intertidal
Intertidal
Intertidal
Intertidal
RMCA MT 1754 (3)
RMCAMT1755(1)
RMCAMT 1748 (1)
RMCAMT1749(1)
RMCA MT 1750 (3)
RMCA MT 1756 (I)
RMCA MT 1751 (1)
RMCAMT 1753 (I)
RMCAMT 1752 (I)
RMCAMT 1747 (1)
SAM A 23250 (1)
SAM UCTECOLCOLL. D 178(1)
SAM MEIRING NAUDECOLL. J 9_1 (6)
SAM UCT ECOL COLL. U 23A (I)
140
JENNIFER M. OLBERS and YVES SAMYN
RESULTS
Details of the collection of different
Ophiocoma species sampled along the KZN
coast of South Africa are presented in Table l.
SYSTEMATIC ACCOUNT
We provide a taxonomic description and
detailed
geographical
distribution
for
Ophiocoma species that are new records for
South Africa. For the other well-known species,
we only provide a summary and cite major
works which provide detailed information on
their taxonomy and distribution.
Order Ophiurida Miiller & Troschel, 1840
Family Ophiocomidae
Ljungman, 1867
Subfamily Ophiocominae
Matsumoto, 1915
Genus Ophiocoma Agassiz, 1836
(Type species Ophiura echinata
Lamarck, 1816 (by subsequent
designation ofClark 1915)).
When distal lobe angular, proximal end
usually more or less truncated to match
interradial separation of adoral shields. Dorsal
arm plates wider than long, fan-shaped, oval
or hexagonal. Ventral arm plates more or less
square-shaped, proximal side straight, distal
side straight to concave. Two tentacle scales,
at least in first arm segments, sometimes only
one towards tip.
Ophiocoma brevipes Peters, 1851
(Plate 1A-G, 2A-C)
Ophiocoma brevipes Peters 1851: 466;
Marktanner-Turneretscher 1887: 303; de Loriol
1893: 25, 26, pl. 23, Fig. 4; Clark 1908: 296;
Clark 1911 : 256; Koehler 1922: 319-322, pl. 72,
Diagnosis
(after Devaney 1970: 9; Clark &
Courtman-Stock 1976: 172-173).
Majority of species large with
D.D. often exceeding 20 mm; three
to seven generally smooth, stout
arm spines, sometimes alternating
three and four on successive
arm segments or on opposite
sides of same segment; lower
arm spines sometimes flattened
and spatulate while upper ones
cylindrical or cigar-shaped. Disc
generally covered with granules,
occasionally concealing scaling on
disc and sometimes extending into
ventral interradial regions. Oral
shields without granules or spines,
triangular with three to four, rarely
five, contiguous oral papillae, outer
one usually widest. Tooth papillae
always present, few to numerous,
with superficial ones in series with
oral papillae. Oral shields large,
oval, hexagonal or pentagonal.
Plate 1. Dorsal (A) and ventral (B) views of Ophiocoma brevipes;
position of juvenile 0. brevipes within the bursal slit of an adult
0. brevipes (C & D); dorsal view of juvenile 0. brevipes (E);
ventral view of juvenile 0. brevipes (F); dental plates (internal and
external view) of adult 0. brevipes (G).
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
141
Material examined
We examined over 150 individuals
of 0. brevipes from South Africa
and Madagascar. On seven of them,
one to three smaller individuals
were attached in such a way that
they had their arms slotted in the
bursal slit. We have here tentatively
treated them as juvenile 0. brevipes.
Adults
D.D.=5.6-25.2
mm;
D.D./
A.L. from 1/3.2 to 1/4.8. Disc
colour patterns variable with
a combination of light greens,
whites, yellows and browns
in blotchy star or simply no
particular pattern. Disc with
small, fine, spherical granules
closely packed on both dorsal
and ventral side. Oral shields
with darker markings, but with
no apparent pattern. Dental plate
(Plate 1G) between 1.9 and 2.1
times longer than wide, with a
wide vertical septum between
each oval, slightly elongated tooth
foramen, dental papillae region
Plate 2. Dental plates of juvenile Ophiocoma brevipes (A); arm limited to approximately 27%
spines of juvenile 0. brevipes (B); distal tip of arms of juvenile 0. of dental plate length. Genital
brevipes (C); dorsal view of Ophiocoma cf. dentata (D); ventral slits clearly visible, elongated
view of 0 . cf. dentata (E); dorsal view of Ophiocoma doederleini and bordered with slightly more
(F); ventral view of 0. doederleini (G) ..
prominent granules. Arms appear
banded (darker) on the dorsal side, some
Figs. 6-9; Devaney 1968: 45; Devaney 1970:
specimens with light dots or specks on ventral
13; Clark & Rowe 1971: 86, 119; Devaney
side of each arm plate along the length of
1974: 151-152; Cherbonnier & Guille 1978:
arm. Ventral arm plates nearly as wide as
168-169, pl. X, Figs. 3,4; Sloan et al. 1979: 104;
long, bluntly pointed on the proximal side.
Clark 1980: 534; Tortonese 1980: 125, Fig. 11;
Dorsal arm plates fan-shaped, much wider
Humphreys 1981 : 10, 23; James 1982: 39-40,
than long. Upper-most spines thickest on the
pl. IlB; Clark 1984: 100; Rowe & Gates 1995:
proximal part of the arm. Longest arm spine
385; Rowe & Richmond 2004: 3292.
less than or equal to the breadth of the dorsal
arm plate. Two tentacle scales.
Location and status of types - ZMB 961 (1
syntype), ZMB 962 (1 syntype), ZMB 963 (1
Juveniles
syntype), ZMB 4660 (3 syntypes).
D.D. = 1.5-4.3 mm; D.D./A.L. from 1/1.8 to
Type locality - Coast of Mozambique (ZMB
1/2.8. Disc colour brown-grey, both dorsally
961-963); Quirimba Island, Mozambique
(Plate lE) and ventrally (Plate 1F). Dorsal
(ZMB 4660).
side of disc marked with five radial pairs
142
JENNIFER M. OLBERS and YVES SAMYN
of bowed brown lines extending from the
margin, passing through radial shields and
almost reaching centre of disc (Plate lE). One
specimen has a faint white dot in the centre of
its disc. Radial shields oval, exposed, minute
in size, hardly visible and similar in colour to
disc. Disc circular in outline and moderately
convex, covered dorsally and ventrally by
imbricated scales of different sizes and shapes.
Disc scales hardly visible dorsally but obvious
ventrally. Disc granulation absent on ventral
and dorsal surfaces. Oral shields and adoral
shields uniformly brownish. Oral shields pearshaped, sometimes with a thickened distal
side, hardly longer than broad. Adoral shields
triangular, almost touching proximally. Jaws
triangular with three to four hyaline-tipped,
somewhat pointed oral papillae on each oral
plate. Dental plate with one to two teeth and
three to four hyaline-tipped dental papillae;
lower-most two to three forming a cluster at
the apex of the jaw angle (Plate IF). Dental
plate (Plate 2A) 1.6 times longer than wide,
with a thin vertical septum between each
oval, slightly elongated tooth foramen, dental
papillae region limited to approximately 28%
of dental plate length. Genital slits unarmed,
almost reaching the margin of disc. Dorsal
arm plates uniformly brown and fan-shaped,
first two to four plates contiguous, thereafter
well-separated. Ventral arm plates uniform
brownish-grey. First two ventral arm plates
slightly broader than long with a straight
distal edge; thereafter plates gradually
become longer, almost rectangular in shape
with the distal edge gradually becoming more
concave. Arm spines light brown with dark
longitudinal line dorsally (Plate 2B). Arm
spines longer than length of arm plates, two on
first segment, three on segments two to five,
thereafter alternating two and three spines
on segments six to nine, and two on further
distal segments, hyaline and ragged structure.
Distally, arm spines with multifid claw-like
hooks (Plate 2C). Single tentacle scale over
whole arm length apart from largest juvenile
(D.D. = 4.3 mm) with two tentacle scales on
first two segments.
Geographical distribution
Tropical Indo-West Pacific region but absent in
the Red Sea, the Persian Gulf and north-western
parts of the Indian Ocean (Rowe & Richmond
2004). Prior to this study, the only record from
southern Africa was Mozambique (Quirimba
Archipelago) (Clark & Rowe 1971). Thus the
present record extends range to Aliwal Shoal,
some 60 km south of Durban, South Africa.
Juveniles were found on adults from South
Africa, Madagascar and Isles Glorieuses.
Ecology
Ophiocoma brevipes is associated with coral
heads or boulders, on fine to coarse sand and
at the base of algal plants in the sandy littoral
zone (between 0-54 m depth, cf. Lane et al.
2000). An overview of known microhabitats of
0. brevipes is presented in Sloan et al. (1979).
On one living adult, the juvenile was
observed to partially move into the genital slit
of the adult after collection (Plate IC, D).
Remarks
The complete absence of granulation on the
disc and the loose meshwork structure of
the dental plate indicate that the specimens
attached to the adult 0. brevipes are most
probably juveniles. The colour pattern
of these juveniles somewhat resembles a
juvenile Ophiomastix venosa which is known
to attach itself to Ophiocoma scolopendrina
(Four'gon et al. 2007). However, based on
the descriptions by Fourgon et al. (2007) and
Cherbonnier and Guille (1978), it can be ruled
out that the nine specimens studied here are
juvenile Ophiomastix venosa, because (i) arm
spines of 0. venosa are distinctly longer, are
glassy and thin; (ii) jaws of 0. venosa are
much more elongated and (iii) ventral arm
plates of 0. venosa taper more proximally.
The proximally extended adoral shields
which nearly meet around the oral shield are
indicative of Clarkcoma Devaney, 1970 and
the alternating number of arm spines, with
the uppermost spine enlarged (approximately
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
two segments long) suggests an affinity with
Devaney's (1970) Scolopendrina Group. On
the other hand, the appressed morphology of the
dental plates is indicative ofDevaney's (1970)
Brevipes Group and, therefore, we believe the
juveniles are 0. brevipes. If this identification
can be confirmed with complementary studies,
such as molecular systematics, this would
present a new case of parental care.
The tips of the arms in the juveniles were
found to bear hooks that may assist in gripping
(see Plate 2C). In addition to the hooks, the
spines have a ragged structure that could also
assist in clinging to the adult (see Plate 2B).
Hooked spines have been also been observed
in ophiuroid species with an epizoic lifestyle
(Hyman 1955).
Ophiocoma cf. dentata Miiller & Troschel, 1842
(Plate 2D, E)
Ophiocoma dentata Muller & Troschel1842 :
99, pl. VII, Figs. 3, 3a; Devaney 1968: 45;
Devaney 1970: 13; Clark & Rowe 1971: 86,
119, pl 18, Figs. 2-3; Cherbonnier & Guille
1978: 168, pl. C, Figs. 3, 4; Tortonese 1980:
125, Figs.llA, B; James 1982: 40, pl. IIC,
D; Rowe & Gates 1995: 386; Price & Rowe
1996: 76; Rowe & Richmond 2004: 3292.
Ophiocoma insularia Lyman 1861: 80-81;
Macnae & Kalk 1958: 130.
Location and status of types - ZMB 931
(holotype, fixed by monotypy ).
Type locality -Unknown according to Muller
and Troschel (1842 : 99). However, the ZMB
has only one specimen of Ophiocoma dentata
in its collection. The catalogue indicates that
this specimen was deposited by Deppe, just
as is indicated by Muller and Troschel (1842),
and that it comes from 'Celebes?' 2, currently
known as the Islands of Sulawesi (Indonesia).
Material examined
D.D. = 14.3 mm, variegated with brown, white
and beige, both dorsally and ventrally with
the presence of small dark brown spots. Oral
2
MUller & Troschell842 give no indication that they
had more than one specimen before them, hence
ICZN Art 73.2.2 applies.
143
shields round, as long as wide, with marbled
pattern. Dental papillae broad, not extending
far into mouth. Dorsal arm plates beige to
brown with a whitish-grey patch surrounded by
dark brown border on the median distal side,
broad and elliptical. Lateral arm plates lighter
with several spots. Ventral arm plates light with
same spots; sometimes a dark coloured patch is
present centrally, square with rounded corners,
as wide as long. Arm spines white to beige,
broadly and irregularly banded once or twice
with light brown. Upper arm spines thick,
blunt, somewhat flattened and slightly shorter
than the lower ones. Tentacle scales, two.
Ecology
According to Devaney (1970), this species
frequents the sub-littoral zone, under boulders
or associated with coral and coral debris on a
sand or rubble substratum.
Geographical distribution
Ophiocoma dentata has a tropical, Indo-West
Pacific distribution (with the exception of the
Red Sea and north-western Indian Ocean (Rowe
& Gates 1995; Rowe & Richmond 2004). Prior
to this study the most southern records originated
from Inhaca Island (Mozambique) (Macnae &
Kalk 1958; as 0. insularia Lyman 1862).
Remarks
This specimen, at first examination, bears
resemblance to Ophiocoma brevipes. Devaney
(1970) pointed out three means by which 0.
brevipes can be separated from 0. dentata
(and 0. doederleini; see hereunder) for
specimens of similar size. First, by comparing
the arm spine sequence: while 0. brevipes
presents five arm spines on segments four to
seven, five to six arm spines up till segment
11, and two to three arm spines thereafter,
0. dentata has only four (occasionally five)
arm spines on segments four to seven, four
arm spines up until segment 13 and three arm
spines thereafter. Secondly, in 0. brevipes the
longest arm spine rarely exceeds the breadth
of the ventral arm plate, while in 0. dentata
(and 0. doederleini) the longest arm spine
greatly exceeds the ventral arm plate. Thirdly,
144
JENNIFER M. OLBERS and YVES SAMYN
by comparing the pigmentation:
while 0. brevipes has a uniform
white or cream colour on the
oral side of arms and the oral
plates and shields, the other two
species have a more grey, brown
or variegated colouration.
However, our specimen
differs somewhat from the
typical Ophiocoma dentata as
reported by Devaney (1970).
First, the arm spine sequence
of our specimen has three
arm spines on its four first
segments, segments five to
fifteen have four arm spines,
and thereafter three arm spines,
which differs from the arms
of 0. dentata where we never
observed five arm spines on
segment seven. Second, the
colouration of the dorsal arm
plates is beige to brown with
a whitish-grey patch that is
bordered by dark brown on
the median distal side, which
differs from the description
given by Devaney (1970,
Fig. 21 ). Finally, the white to
beige arm spines are broadly
and irregularly annulated with
light brown. Given these three
differences, we are reluctant to
describe the single specimen
as a new species until more
material becomes available.
Ophiocoma doederleini de
Lorio1, 1899
(Plate 2F, G and Plate 3A, B)
Ophiocoma doederleini de Loriol
1899: 30, pi. 3, Fig. 2; Devaney
1968: 69; Devaney 1970: 1218, Figs. 18, 14, 22, Table 2B,
3; Devaney 1974: 154; Sloan et
al. 1979: 104, Figs. 8-10; Clark
1980: 534; Humphreys 1981: 10,
24; Clark 1984: 100; Rowe &
Gates 1995: 396.
Plate 3. Dorsal views of reticulated (A) and spotted (B) forms of
Ophiocoma doederleini; dorsal view of Ophiocoma erinaceus (C);
dorsal view of juvenile 0. erinaceus (D); ventral (E) and dorsal (F)
views of Ophiocoma pica (from Mahe, Seychelles).
Ophiocoma dentata Lutken 1859: 165 (non Muller &
Troschel1842); Clark 1921: 121.
Location and status of types - MHNG INVE 71892
(Holotype).
Type locality- Mauritius.
Material examined
The specimens collected represent the two known colour
forms (Devaney 1970; Sloan et al. 1979).
Whole specimens greyish-brown dorsally and ventrally
(Plate 2F, G). Disc greyish-brown with fine black reticulating
lines, white-ringed black spots, or speckled with light spots.
Spots, speckles and lines do not outline the shape of radial
shields, thus radial shields are not conspicuous (Plate 3A,
B). Oral shields large, round to oval in shape, white on edges
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
and often with large irregular grey blotches.
Dental papillae broad but, lower-most shorter
than others. Ventral side of arms brown with
spotted white bands or with dark bands, with
narrower bands in between which continue
down the arm. Arm spines taper, annulated
white and/or grey. Combination of two and
three tentacle scales' on arms.
Ecology
All specimens were present under large
boulders over gravel. Rowe and Gates (1995)
record it as a benthic, inshore, littoral species.
An overview of known microhabitats of
Ophiocoma doederleini is presented in Sloan
et al. (1979).
Geographical distribution
Indian Ocean and west central Pacific Ocean
(Rowe & Gates 1995). Our specimens
represent a new record for southern Africa.
Remarks
Devaney (1970: 15-16, Table 2b) provides
an accurate means to separate Ophiocoma
dentata from 0. doederleini: annulation
of the arm spines is easily used in the field
and is absent in 0. dentata but present in 0.
doederleini.
Ophiocoma erinaceus Miiller & Troschel, 1842
(Plate 3C, D)
Ophiocoma erinaceus Muller & Troschel
1842: 98; Devaney 1968: 173 (synonymy3);
Devaney 1970: 33, figs 45-47; Kalka 1958:
207, 216, 237; Clark 1967: 47; Clark &
Rowe 1971:86, 119, pi. 17, figs 5, 6; Clark &
Courtman-Stock 1976: 122, 173; Cherbonnier
& Guille 1978: 169, pl. X, Figs. 5, 6; Sloan et
al. 1979: 106, Figs. 11, 12; Clark 1980: 535,
548; Tortonese 1980: 124; Humphreys 1981:
10, 24; James 1982: 38, pi ID; Price 1982: 8;
Clark 1984: 100; Rowe & Gates 1995 : 387;
Price & Rowe 1996: 77; Rowe & Richmond
3
Cherbonnier & Guille (1978) removed 0.
schoenleinii Miiller & Troschel, 1842 - characterized
by the presence of just one tentacle scale, not two
as in 0. erinaceus- from Devaney's synonymy. We
follow Cherbonnier & Guille (1978).
145
2004: 3292; O'Hara et al. 2004: 537-541;
Benavides-Serrato & O'Hara 2008: 51; Reza
Fatemi et al. 2010: 44, Fig. 2.
Ophiocoma similanensis Bussarawit & Rowe
1985: 1, Figs. 1, 2; Price and Rowe 1996: 77.
Location and status of types - ZMB 921
(Syntype 1); ZMB 922 (Syntype 2); ZMB
923 (Syntype 3), specimen lost; ZMB 924
(Syntype 4), specimen lost.
Type locality - Red Sea and Indian Ocean.
Material examined
D.D.
2.8-21.5
mm.
Specimens
characteristically black, dark brown or dark
red dorsally and lighter ventrally. Some of
the specimens under study were juveniles as
evident from pigmentation of the dorsal disc
(pairs of cream lines starting at the margin,
passing through the radial shields and almost
reaching the centre of the disc; Plate 3D), the
arm spines (edge lighter) and the armament
of the disc (disc dorsally and ventrally devoid
of granules) (see also Cherbonnier & Guille
1978: 171; Bussarawit & Rowe 1985, as
Ophiocoma similanensis). Oral shields pearshaped, distally broadest. Dorsal arm plates
uniform black, fan-shaped, distally convex,
overlapping as tiles on a roof, more than twice
as wide as long. Ventral arm plates uniform
brown, from regular hexagons proximally to
pentagons distally. Two equal-sized tentacle
scales over the complete arm. Three to four
arm spines, with uppermost one always
largest. Three specimens had longitudinal
stripes on arms spines similar to the juveniles
of this species. Tube feet of the live specimens
were reddish. Arm spines on most specimens
flattened close to disc.
Ecology
Benthic, inshore (Rowe & Gates 1995) from
0-27 m depth (Lane et al. 2000). They are
associated with coral (Clark & CourtmanStock 1976; Humphreys 1981; Bussarawit &
Rowe 1985; StOhr et al. 2008) and are often
found on gravel under boulders. Juveniles
were found on an encrusting turret sponge
(Haliclona sp.) or under dead coral boulders.
146
JENNIFER M. OLBERS and YVES SAMYN
An overview of known microhabitats of
Ophiocoma erinaceus is presented by Sloan
et al. (1979).
Geographical distribution
Tropical to subtropical Indo-Pacific region.
Remarks
Even though 0. erinaceus is one of the
most abundant and conspicuous brittle stars
in littoral tropical seas, its taxonomy has
only recently been elaborated. O'Hara et al.
(2004) used molecular, morphological and
day/night colour change data to show that
0. erinaceus is a species complex of three
species: 0. erinaceus, 0. schoenleinii Muller
& Troschel, 1842 and a third undescribed
species. The latter was formally described
and named 0. cynthiae by Benavides-Serrato
and O'Hara in 2008. Species in the complex
can be distinguished from one another by: (i)
colouration of tube feet (red in live or white
in preserved 0. erinaceus; grey in life and
preserved 0. cynthiae and 0. schoenleinii),
(ii) number of tentacle scales (one in 0.
schoenleinii; two in 0. erinaceus and 0.
cynthiae); (iii) granulation of the ventral side
of the disc (largely absent in 0. cynthiae, as
a wedge near the margin in 0. schoenleinii
and extending almost to the oral shields in 0.
erinaceus); and (iv) the size and morphology
of the dental plates. Price and Rowe (1996)
recognised that their 0. similanensis
Bussarawit & Rowe, 1985 is but a juvenile 0.
erinaceus and described growth changes for
specimens ranging from a D.D. of 3.6-22.2
mm. The juveniles in this study match the
description of 0. similanensis very well.
Ophiocoma pica Muller & Troschel, 1842
(Plate 3E, F)
Ophiocoma pica Muller & Troschel 1842:
101; Clark 1921: 127, pl. 13, Fig. 8; Clark
1938: 333; Balinsky 1957: 25-26; Devaney
1968: 131; Macnae & Kalk 1958: 130;
Devaney 1970: 19, Figs. 23 & 24, 20, Figs.
27, 25; Clark & Rowe 1971 : 86-87, 118; Clark
& Courtman-Stock 197 6: 173; Cherbonnier &
Guille 1978: 172, pl. XI, Figs. 5, 6; Sloan et
al. 1979: 106, Clark 1980:535, 548; Tortonese
1980: 124; Price 1982: 8; Clark 1984: 100;
James 1982: 36-38, pl IC; Rowe & Gates
1995: 387; Price & Rowe 1996: 77.
Location and status of types - Unknown;
placed in MNHN according to Muller and
Troschel (1842: 101) and according to the
MNHN catalogue but not in MNHN according
to Dr N. Ameziane (pers. comm.).
Type locality -Unknown according to Muller
and Troschel (1842). The senior subjective
synonym (Lyman 1865) Ophiocoma lineolata
Desjardins in Muller and Troschell842 stems
from Mauritius.
Material examined
D.D. = 5.3 mm (D.D./A.L = 1/4). Disc covered
dorsally with spherical granules extending
onto distal parts of ventral interradii. Colour,
pale yellow (after preservation in alcohol). Oral
shields mainly oval; adoral shields triangular,
not contiguous proximally; oral papillae three
to four, dental papillae six to ten; teeth one
or two, slightly elongated and blunt. Genital
papillae present, cone-shaped. Dorsal arm
plates fan-shaped, convex on distal side and
concave on proximal side, hardly changing
shape distally. Ventral arm plates straight
to slightly convex on distal side, concave
proximally, plates becoming slightly longer
distally. Arm spines five proximally and four
distally, slender, second spine longest, about
two times segment length, lower arm spines
same length as segment or slightly longer.
Tentacle scales two, inner one slightly smaller
proximally.
Ecology
Benthic, inshore (Rowe & Gates 1995), 0-24
m (Lane et al. 2000). Usually associated with
coral (Clark 1952; Devaney 1968, 1970;
Clark & Courtman-Stock 1976; Price &
Rowe 1996) but also found under rock or dead
coral rubble (Devaney 1970). An overview
of known microhabitats of Ophiocoma pica
is presented by Sloan et al. (1979).
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
147
Geographical
distribution
Widely distributed throughout
Indo-Pacific region (Clark 1921;
Clark & Rowe 1971).
Remarks
Given the material from South
Africa was in a poor state,
we chose to illustrate (Plate
3E, F) a specimen from Mahe
(Seychelles).
The
record
discussed here presents a range
extension from Richards Bay to
Qolora (Eastern Cape Province).
Colour in life dark brown or
black with radiating golden lines
on disc and, often, transverse
bands annulating the arms.
Ophiocoma pusilla (Brock,
1888) (Plate 4A, B)
Ophiomastix pusilla Brock
1888: 499; Devaney 1970: 25
(records before 1970).
Ophiocoma
latilanxa
Murakami 1943a: 194-196;
Murakami 1943b: 218; Devaney
1970: 25-27.
Ophiocoma pusilla (Brock,
1888); Clark 1921:
131;
Devaney 1970: 25, Figs. 26, 29;
Cl ark & Rowe 1971: 86-87, 118;
Clark & Courtman-Stock 1976:
122, 174, Fig. 190; Cherbonnier
& Guille 1978: 173-174, pl. XI,
Figs. 3,4; Sloanetal. 1979: 106;
Clark 1980: 535, 544; Tortonese
1980: 127; Humphreys 1981:
10, 24; Price 1982: 8; Clark
1984: 100; Rowe & Gates 1995:
388; Price & Rowe 1996: 77.
Location and status of types ZMB 5429 (Lectotype); ZMB
4777 (Paralectotype).
Type locality
Indonesia.
Ambon,
Plate 4. Dorsal (A) and ventral (B) views of Ophiocoma pusilla;
dorsal (C) and ventral (D) views of Ophiocoma scolopendrina
(Neotype: MNHN EcOh 11043); dorsal (E) and ventral (F) views
of Ophiocoma valenciae.
Material examined
D.D. = 3.3-7.7 mm. Disc of one specimen slightly speckled
while another specimen had banded arms from halfway down
the arms to the tips. Dorsal disc with uniformly distributed
granules concealing radial shields. Ventral disc with same
type of granules, leaving bare only a narrow V-shaped
interbrachial area. Oral shields oval, nearly twice as long as
wide. Adoral shields triangular, not touching proximally. Four
to five oral papillae per jaw. Dental papillae in two to three
rows. Dorsal arm plates proximally fan-shaped, wider than
long, with convex distal side touching the next plate only for
about a third of its width; distally plates longer than wide and
less contiguous. Ventral arm plates fan-shaped, broader than
long, though distally becoming longer than broad. Four to five
arm spines, hollow, glassy and about 2.5 times the segment
length. Second uppermost arm spines at a third of arm length
JENN1FER M. OLBERS and YVES SAMYN
148
with pustular distal expansions, while other
arm spines taper (cf. also Clark & Rowe 1971 ).
Tentacle scales, two.
Ecology
Benthic, inshore (Rowe & Gates 1995), 0-20
m depth according to Lane et al. (2000), our
deepest specimen was found at 32 m depth.
Known to occur in sand channels, under rubble
and associated with coral (Clark & CourtmanStock 1976; Humphreys 1981 ; Price & Rowe
1996). An overview of known microhabitats
of Ophiocoma pusilla is presented by Sloan
et al. (1979).
Geographical distribution
Ophiocoma pusilla has a tropical Indo-West
Central Pacific Ocean distribution (Rowe &
Gates 1995), including the Red Sea (Clark
1967, as Ophiomastix pusilla, Price 1982). In
southern Africa, this species was reported from
Mozambique (Clark & Courtman-Stock 1976).
Remarks
After examination of the size, shape and
sequence of the arm spines, the nature of the
dental plates and oral shields and the number
of dental papillae, Devaney (1970) concluded
that Ophiocoma latilanxa Murakami, 1943,
is a junior synonym of 0. pusilla. In 1989,
Soliman ignored this when he identified 0.
latilanxa4 from the Red Sea. In 1991, a new
species, Ophiocoma aegyptiaca Soliman,
1991, was described from the same area
and Soliman (1991) continued to treat 0.
latilanxa as a valid species and noted that his
new species bears close resemblance to 0.
schoenleinii and 0. latilanxa (= 0. pusilla) .
Although we have not seen the type material,
we suspect that 0. aegyptiaca and 0. latilanxa
(= 0. pusilla) will prove to be synonyms.
According to Soliman ( 1991 ), the differences
between 0. aegyptiaca and 0. latilanxa
consist mainly of: (i) the colour pattern of
the disc and the plates (our specimens of
0. pusilla show that the colouration is very
4
Soliman 1989 misspelled Ophiocoma latilanxa as 0.
latilanaxa.
variable (disc uniform brown to reticulate to
spotted with dark blotches; spines uniformly
coloured to spotted)); (ii) the shape of the
oral shields (although Soliman ( 1991) states
that the oral shields of 0. aegyptiaca are
trapezoidal he drew them oval (Fig. 3A, p. 82
& p. 85), similar to the ones of 0. pusilla);
(iii) the number of arm spines (maximum five
in 0. aegyptiaca, except Soliman (1991) does
not give the sequence of the arm spines; our
0. pusilla specimens mostly have arm spine
sequences: 3-3-4-4-4( or 5)-5-4-5). On the
other hand the form and size of the dorsal arm
plates are very similar for 0. aegyptiaca and
0. pusilla (compare Devaney 1970: Fig. 29 p.
21 with Soliman 1991 : Fig. 4 p. 83).
Ophiocoma scolopendrina (Lamarck, 1816)
(Plate 4C, D)
Ophiura scolopendrina Lamarck, 1816: 544.
Ophiocoma scolopendrina (Lamarck, 1816):
Kalk 1958: 205 ; Macnae and Kalk 1958: 130;
Devaney 1968: 203; Devaney 1970: 33-35;
Clark & Rowe 1971: 86, 119, pl. 17, Figs. 3,
4; Clark & Courtrnan-Stock 1976: 122, 174;
Sloan et al. 1979: 106, Fig. 13; Clark 1980:
535; Tortonese 1980: 124; Price 1982: 8;
James 1982: 36-39, pl. IIA; Rowe & Gates
1995: 388; Reza Fatemi et al. 2010: 45, Fig. 3.
Location and status of types - Unknown for
0. scolopendrina, not in the MNHN (Dr M.
Eleaume, pers. comm.). According to Devaney
(1968), type specimens are present in the Dorpat
Museum as the junior subjective synonym
0. variabilis Grube, 1857 (from 'Waohu
Island'- Oahu Island?) and in the Museum of
Comparative Zoology as the junior subjective
synonym 0. molaris Lyman, 1861 (from
'Kingsmill Islands' = Gilbert Islands, Kiribati).
Type locality - Recorded by Lamarck (1816)
as Mauritius ('lie de France'). The ZMB
holds two syntypes of a junior synonym, 0.
alternans von Martens, 1870, from Java.
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
Material examined
Given that current synonymy might lead to
some confusion in its distribution, we felt it
appropriate to establish a neotype from the
original type locality, Mauritius.
Type material
N eo type Ophiocoma scolopendrina (Lamarck,
1816), here designated: MNHN EcOh 11043
(specimen with D.D. = 23.8 mm), Mauritius,
coil. M. Carie, 1913.
Disc uniformly brown both dorsally and
ventrally, although where granules have been
worn off, lighter patches visible. Dorsal arm
plates blotched with brown on beige, giving
arms a variegated to banded pattern. Oral
and adoral shields with similar blotching.
Arm spines uniform in colour (light brown
ventrally and somewhat darker dorsally),
although on rare occasions some banding
can be observed on uppermost spines. Disc
pentagonal with interradial margins straight to
slightly indented. Dorsal disc densely covered
with spherical granules, covering the whole
surface including the radial shields which
cannot be distinguished. Ventral disc with
same, densely distributed granules, but less
dense closer to genital slit which is bordered
by a fringe of elongated genital papillae. Oral
shields oval, shorter (2.5 mm) than wide (3 .0
mm). Adoral shields restricted to the lateral
edge of the oral shield, triangular but with other
margin curved, not touching proximally. Five
oral papillae on each oral plate, inner ones are
more pointed than outer, wider ones; buccal
tentacle scale very low and wide. Four to nine
dental papillae, placed in a cluster below wide
truncated teeth. Dorsal arm plates fan-shaped,
wider than long with distal margin straight
in first segments, becoming more convex in
distal segments, plates contiguous throughout
the arm. First two ventral arm plates distinctly
smaller than rest, with distal margin indented,
lateral margins convex and proximal margin
straight, about as long as wide. Subsequent
ventral arm plates significantly larger, wider
then long, with convex distal margin and
concave proximal margin which is only
slightly overlain by preceding plate, laterally
149
plates recurved around tentacle pore. Arm
spines three to five (three on segment three,
four to five on segment eight), uppermost
ones thick, short, but longer than dorsal arm
plates; lower arm spines more slender, always
longer than dorsal arm plates, except for first
two segments. Arm spines uniform in colour
(light brown ventrally and somewhat darker
dorsally), although on rare occasion some
banding can be observed on uppermost spines.
Two oval, tentacle scales, inner one a fraction
longer, over the complete arm length.
Non-type material
To avoid damage to the neotype, we used
another specimen to describe the dental plate
which is between 1.9 times longer than wide,
with a very wide vertical septurn between
each oval, elongated tooth foramen; dental
papillae region limited to approximately 20%
of dental plate length.
Remarks
As noted by Devaney (1968) and others,
0. scolopendrina is often confused with 0.
macroplaca (Clark, 1915), a Hawaiian endemic,
and 0. erinaceus Muller & Troschel, 1842. We
did not have 0. macroplaca specimens at our
disposal so will refrain from commenting on
Devaney's (1970) means of distinguishing it from
0. erinaceus and 0. scolopendrina. On the other
hand, our limited comparative study of the dental
plates of 0. erinaceus, and 0. scolopendrina
shows that dental plate morphology can be used
to recognize both species with certainty.
The
senior
primary
homonym,
Ophiocoma scolopendrina Agassiz, 1836 (as
for instance mentioned by Muller & Troschel
1842), is to be considered a nomen nudum.
Ecology
Benthic, inshore, littoral (Rowe & Gates
1995), 0-13 m depth (Lane et al. 2000).
An overview of known microhabitats of
Ophiocoma scolopendrina is presented by
Sloan et al. (1979).
150
JENNIFER M. OLBERS and YVES SAMYN
Geographical distribution
DISCUSSION
Tropical, ludo-Pacific region (Rowe & Gates
1995), including the Red Sea.
A total of 70 Ophiocoma specimens
belonging to eight species were collected
during the five expeditions to KwaZuluNatal. 0. brevipes was the most common
(n=24) followed by 0. pusilla (n=l4) and 0.
valenciae (n= l4). Only a single individual
of 0. dentata was collected in South Africa.
Although previously recorded in the Eastern
Cape, no 0. pica or 0. scolopendrina were
collected by us. 0. brevipes, 0. doederleini,
0. pusilla and 0. dentata are new records for
South Africa. Even though Martynov (2010)
warned that dental plates are to be used with
caution as a taxonomic character, the dental
plates permitted us to assign the juveniles in
our study to the Brevipes Group, as suggested
by Devaney (1970).
According to Hendler (1975), only 55
species of ophiuroids have been reported as
viviparous, which is less than 3% of all known
species. If the juveniles we found attached to
adult 0. brevipes individuals are indeed also
0. brevipes individuals, this report is the first
account of brooding behaviour and parental
care in 0. brevipes. However, based on the
examined material, we conclude that freeliving juveniles of 0. brevipes must be very
rare, which is in contrast to other ophiocomid
species such as 0. erinaceus which have been
found as free-living (Price & Rowe 1996).
Reproductive experiments with 0. brevipes
populations from various locations will
probably provide an insight into the ontology
and the reproductive strategy of 0. brevipes.
Ophiocoma valenciae Muller & Troschel,
1842 (Plate 4E, F)
Ophiocoma valenciae Muller &
Troschel 1842: 102; Devaney 1968: 126;
Eyre & Stephenson 1938: 38, 43; Kalk
1958: 200, 207, 237; Macnae & Kalk
1958: 130; Clark 1967 : 44-45; Macnae &
Kalk 1969 : 101, 106, 130; Clark & Rowe
1971: 86, 119, pi. 18 Fig. 1; Sloan et al.
1979: 109, Fig. 14; Clark 1980: 535, 548;
Tortonese 1980: 125; Humphreys 1981 :
10, 24-25; Price 1982: 8.
Location and status of types - ZMB 4625
(Syntype 1); ZMB 955 (Syntype 2).
Type locality - Aden.
Material examined
D.D. = 7.7-20.3 mm. Disc colour brownish,
arms tawny with darker bands; one specimen
lacking darker bands on arms. Disc covered
dorsally and ventrally with moderately fine
granules, which become elongated towards
margin of disc. Radial shields defined by
lighter colour on some specimens, but could
be an artefact due to preservation. Dorsal arm
plates broad, oval with up to six arm spines
proximally. Uppermost arm spines shorter
than middle spines, as long as arm width.
Tentacle scale, one.
Ecology
Associated with coral (Day 1969) and
sponges (Humphreys 1981). Found within
rocky crevices, cobbles, rubble and various
algal beds (Humphreys 1981). An overview
of known microhabitats of 0. valenciae is
presented by Sloan et al. (1979).
Geographical distribution
Tropical Indian Ocean, including Red Sea and
possibly the Persian Gulf (Clark and Rowe
1971; Tortonese 1980).
Acknowledgments-It is with great pleasure that
we thank Dr F. Rowe (Senior Research Fellow,
Australian Museum, Sydney, Australia) for
reading and commenting on the first drafts of
this paper. We are also greatly in debt to Dr D.
VandenSpiegel who gave us access to collections
in the RMCA and who assisted in documenting
the specimens. Dr C. Lueter (ZMB), Dr N.
Ameziane and Dr M. Eleaume (MNHN), Dr J.
Mariaux (MHNG) and Ms S. Halsey (NHM)
provided information on the ophiocomid types
they curate. We also express our appreciation to
Ms K. Vrancken (RBINS) who re-worked some
The Ophiocoma species (Ophiurida: Ophiocomidae) of South Africa
of the figures. The Belgian National Focal Point
to the Global Taxonomy Initiative is wannly
thanked for financial and logistical support to
both authors. The comments of Dr T. O'Hara
and an anonymous reviewer greatly improved
the paper.
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